the Host Range of Bactra truculenta Meyrick and

Vol. XVIII, No. 3, June, 1964 417 Observations on Nutgrass Insects in Hawaii with Notes on the Host Range of Bactra truculenta Meyrick and Athesapeuta cyperi Marshall George O. Poinar, Jr.1 DEPARTMENT OF BIOLOGICAL CONTROL UNIVERSITY OF CALIFORNIA CITRUS RESEARCH CENTER AND AGRICULTURE EXPERIMENT STATION RIVERSIDE {Submitted for publication October, 1963) INTRODUCTION Throughout many parts of the world, the purple nutgrass, Cyperus rotundus L., is regarded as a troublesome weed and often as an important pest. A closely related species, C. esculentus L., frequently called yellow nutgrass or chufa, is also often classified as a weed, although in certain areas it is a source of food for wildfowl, livestock and occasionally even for humans. Both species are believed to have originated in the Mediterranean region, and various control practices have been employed throughout the world for their destruction. Because of its more durable rhizome system, purple nutgrass has resisted both cultural and chemical control measures in many areas and has been investigated for possibilities of eradication by means of biological control. Hawaiian scientists were the first to attempt this when they studied insect pests of nutgrass in the Philippines and introduced two insects from that country into Hawaii in 1925 to combat C. rotundus (Swezey 1926, Pemberton 1948). These two insects were Bactra truculenta Meyrick (Olethreutidae) and Athesa peuta cyperi Marshall (Curculionidae) which soon became established on Oahu and spread to the islands of Hawaii and Maui. Initially, the moth seemed most promising (Swezey, 1927-1928), but eventually native parasites reduced its effectiveness (Swezey 1931, 1935, 1946). The weevil was somewhat slower in spreading, but now, at least on several of the islands, has a range equal to that of B. truculenta (Swezey and Pemberton, 1931). The author spent three and one-half months (April 15-July 31, 1963) on the island of Hawaii studying the host range of B. truculenta and A. cyperi to determine if introductions into southern California for biological control of nutgrass would be safe. During these studies, there was ample opportunity to examine the insect fauna of both C. rotundus and C. esculentus. 1 Grateful acknowledgement is made to Clifton J. Davis and Ernest R. Yoshioka, who supplied laboratory space and facilities for these studies at Hilo, Hawaii, and to F. A. Bianchi, N. L. H. Krauss, and J. W. Beardsley, whose help in locating available literature on the flora and fauna of the islands is deeply appreciated.

418 Proceedings, Hawaiian Entomological Society BACTRA TRUCULENTA MEYRICK Life history studies: This olethreutid was first described by Meyrick from India (199). It is also recorded from the Philippines and Burma and may possibly occur in South America (Williams, 1931). Both Ghosh (1921) and Williams (1931) have published brief accounts on the biology of this moth and further findings by the author follow below. The eggs, laid either in the midrib groove or on the smooth surface of the nutgrass leaf, are flattened and usually overlapping in a row. Several hundred eggs were held at an average temperature of 74.2 F. (range of 65.6-82.8) over a two-week period and found to hatch in Rye and one-half to six days at an elevation of 75 feet. The eggs are white when first laid, and the embryo can be distinguished after two to three days. The eyes are visible after three to four days and the internal organs in five to six days. At the time of hatch, the larva is pale yellow with a black prothoracic shield and head capsule. Just before emergence from the egg, the larva initiates biting motions against the chorion. As this behavior becomes more intensified, the head is swung back and forth as the larva searches for other possible emergence areas. Eventually, the chorion is broken and the larva makes its exit. After emergence, many larvae crawl a short distance away from the egg, burrow into the leaf and tunnel toward the base of the plant. They may later emerge from the leaf and start a new tunnel or simply crawl over the plant and eventually make their way down to the base of the stem where they tunnel directly into the stem tissue. A few of the larvae drop off the leaves, suspended by a delicate silk strand, and are blown to neighboring plants. After entering the plant, the larvae make their way to the meristematic stem tissue and some eventually reach the bulb and feed on its contents. The first sign of infestation is the yellowing of the inner leaves of the plant as they are severed from the main portion of the stem. These leaves wither and die and usually the whole plant eventually succumbs. Sometimes an adventitious shoot is sent up from the base of the plant which survives after the parent plant dies. Under laboratory conditions, the larval period lasted from 18 to 2 days at an average temperature of 75 F. The pupa was formed in a lightly constructed cocoon in the basal stem portion of the plant. The adult moth emerged in about a week and was capable of ovipositing in about two days. Two females that were reared from larvae were caged until death and found to have laid 77 and 13 eggs, respectively. Parasites: The gradual lack of effectiveness oibactra on nutgrass in succeeding years after its introduction was generally attributed to parasitization of the eggs by Trichogramma minutum (Swezey 1931, 1931a, 1946). The author found egg clusters of Bactra which ranged from to 1 percent parasitized by T. minutum. Parasitized eggs appear totally black to the naked eye, in contrast to the yellow color of unparasitized eggs. This color appears to be due to a darkening of the inside of the chorion and not to the body of the parasite itself. The only other recorded parasite of B. truculenta appears to be Chelonus blackburni Cameron (Braconidae) which was recorded from one larva on the-

Vol. XVIII, No. 3, June, 1964 419 island of Maui (Swezey 1929). However, I discovered that during May and June many of the moth larvae feeding on yellow nutgrass in the sugar plantations were ^--parasitized by two ichneumonids, Diadema blackburni (Cameron) and Eriborus sp.2 Each is a larval parasite which emerges from the body of the caterpillar after it has spun its cocoon and then forms its own cocoon inside the Bactra cocoon. Host plant studies: Very little information is available concerning the host range of Bactra tmculenta. Pemberton (1948) mentions that tests were made with this moth in Honolulu in 1922 on economic plants, but I have been unable to find any account of these experiments. According to all previous records, it appeared that B. truculenta was confined to Cyperus rotundus and substantiating this was the report that it had never been recorded injuring any crop or garden plant in India or the Philippines (Swezey, 1926). Since its introduction into Hawaii, however, it has been discovered feeding on Kyllingia monocephala Rottb. (Swezey 1929) and K. pumila Michaux (Williams, 1931). In order to insure that this insect would not harm other plants if introduced into California, host plants studies were conducted with plants in the Gramineae, Cyperaceae, and Juncaceae. The results are shown in table 1. In the oviposition studies, 1 to 15 adult moths were caged with the host plant. In the larval feeding tests, newly hatched larvae were placed both on and in potted and cut plants. Results indicate that the adults will oviposit on a wide variety of plants; however, the feeding habits of the young larvae are much more restricted and of the plants tested, development can be completed only on C. rotundus, C. esculentus, C. alternifolius L. and C. papyrus L. Only a few larvae were able to com plete their development on the latter two plants, and the pupae were all destroyed by ants before they could mature. I have been unable to locate infested plants of C. papyrus and C. alternifolius in the field, but Bactra larvae are quite common in esculentus. In order to determine if Bactra adults showed any preference in regard to oviposition on various host plants, a number of potted plants were trimmed to about the same surface area of foliage and placed in a large screen cage with 2 adult moths. After 4 days, the plants were removed and examined for eggs. The number of eggs laid on each plant is as follows: CyperusrotundusL 37 Sorghum halepense (L.) Pers 29 Cyperus esculentusl 17 Cyperus alternifolius L 17 Avena sativa L 9 Tropaeolum majus L. (nasturtium) The results indicate that the adults do show a preference, yet plants other than the desired host (C. rotundus) are also chosen for oviposition. The first instar of Bactra truculenta was unable to live on any of the other sedges and the rushes and grasses tested. When later instars were placed on corn, wheat, and oats, they fed for a day or two and then succumbed. 2 Determined by Dr. L. M. Walkley, Entomology Research Division, Beltsville, Maryland.

42 Proceedings, Hawaiian Entomological Society ATHESAPEUTA CYPERI MARSHALL Life history studies: This barine was described by Marshall in 1928 and is known to occur only in the Philippines. Close relatives are found in India (Marshall, 1928) and Burma (Ghosh, 1921). Table. 1. Results of Host Plant Studies for Ovipositional Sites and Larval Feeding of Bactra truculenta Meyrick Host Plants Eggs Deposited Larvae Fed (first instar) Development Completed Gramineae Avena sativa L Triticum vulgare Vill Bromus unioloides H.B.K Sorghum halepense (L.) Pers Zea mays L Festuca elatior L Poa annua L Hordeum vulgare L Lolium perenne L Dactyles glomerata L Cyperaceae Cyperus esculentus L C alternifolius L C. papyrus L Carex wahuensis C. A. Meyer var. rubiginosa R. Krauss Eleocharis obtusa Schultes Scirpus lacustris L Fimbristylis polymorpha Boeckeler Killingia pumtla Michaux Juncaceae Luzula campestris (L.) DC. var. hawatiensis (Buch.) Degener and Fosberg Juncus bufonius L +++I++++++ +OOOO +Illlllllll +1111 II + + Plant died =not tested + =yes As with most of the weevils of this tribe, little of their biology is known. The following observations will supplement Williams' notes (1931) on the biology of this weevil. Adults were kept in petri dishes and a fresh portion of the nutlet, stem, and lower portion of nutgrass leaves was supplied every other day. Eggs were deposited in cavities in the tubers of the plants. They were about one-fourth mm. long, pearly white when laid, and remained so except for the head capsule which turned black just before hatching. The incubation period lasted four to five days at an average temperature of 75 F. The young larvae apparently enter the base of the stem and make their way to the meristematic tissue and down into the tuber where they feed on its contents.

II 1o1++ o1o1 Vol. XVIII, No. 3, June, 1964 421 Pupation occurs in the hollowed-out tuber, and the adult is formed after 1 to 11 days (at average temperature of 75 F.). The newly formed adults seem to spend days inside the tuber before making their way to the outside. In the field during the day, the adults could be found in the soil, hidden between the bases of the leaves, or rarely exposed on the leaf surface. In the laboratory, they fed on all parts of the plant but preferred the basal stem portion and the nutlet. The adults appear to have a preoviposition period which may last up to two months or longer. Mating was observed one month after emergence, but oviposition occurred much later. No parasites of the weevil were recovered. Host plant studies: Because of the long preoviposition period of the nutgrass weevil, there was no opportunity to test various plants for oviposition sites and determine the host range for the early larval instars. Instead, adults were placed on various plants and third and fourth larval instars were transferred from nutgrass stems obtained in the field and placed on stems of various test plants in the laboratory. The results are shown in table 2. In the laboratory, the adults fed only on Table 2. Results of Host Plant Tests with Adults' and Older Larvae of Athesapeuta cyperi Marshall Host Plants Larval Stage Larvae Fed Development Completed Adult Fed Gramineae Avena sativa L Triticum vulgare Vill Zea mays L Triticum vulgare Vill Cyperaceae Cyperus papyrus L C. esculentus L C. alternifolius L Carex wahuensis Meyer var. rubiginosa Krauss. Juncaceae Juticus bufonius L Other Bidens sp Tropaeolum majus L. (Nasturtium) Coreopsis sp 3rd 3rd 3rd 3rd and = not tested + =yes -=no C. esculentus L. and C papyrus L. The data gained from the larval trials suggest that, as with the moth, later larval instars can complete the remainder of their development on a wider range of plants than can the first instars. In the field, C. esculentus was heavily attacked and appears as acceptable to the weevils as C. rotundus. Larvae were also found in C. polystachyos Rottb., and an adult was observed feeding on K. pumila.

422 Proceedings, Hawaiian Entomological Society FIELD OBSERVATIONS In general, both insects appear to be more plentiful at the lower elevations in Hawaii although the weevil has been recorded as high as 3, feet on Oahu (Zimmerman, 1936). Both the moth and the weevil are often found in the same locality and although they may occur side by side, have never been found together in one plant. In fact, I have never found more than one of these insects in a single nutgrass plant. In plants of C. esculentus in the sugarcane field on Hawaii, the weevil appeared more abundant than the moth in May, and out of 25 plants examined, six were infested with the weevil and three with the moth. This may be due to parasitization of the moth, however. In a macadamia nut orchard infested with C. rotundus, the moths were much more plentiful than the weevil and continued breeding throughout the summer months. It is difficult to say what factors play the leading role in the abundance of these two insects. The extremes of temperature and moisture found in Hawaii offer a good chance to study the effect of physical factors on the distribution of these insects. REPORTS OF HAWAIIAN NUTGRASS INSECTS A number of other insects have been reported feeding on C. rotundus in Hawaii at one time or another and will be briefly discussed below. Most of these are not specific to nutgrass. Two species of mealybugs, Dysmicoccus brevipes (Cockerell) and Geococcus radicum Green have been reported on the roots of nutgrass, C. rotundus, in the Hawaiian Islands. Both also attack pineapple. Two species of thrips, Taeniothrips cypraceae Bianchi and Haplothrips (Hindsiana) sakimurai Moulton have also been reported attacking C. rotundus in Hawaii. Both are found on other plants. The aphid Schizaphis cyperi (Van der Goot) was first found in the Hawaiian Islands in 1939 and feeds on the above-ground portion of C. rotundus L. (Zimmer man, 1948). Several species of Lepidoptera have been reported feeding on C. rotundus in Hawaii. These are Spodoptera mauritia Boisduval3 (Williams, 1931), Spodoptera exempta (Zimmerman, 1958), and Pseudaletia unipuncta (Zimmerman, 1958). Beardsley (1963) noted adults of Sphenophorus cariosus (Olivier) around the roots of nutgrass, C. rotundus, and observed adults feeding and ovipositing on this plant under laboratory conditions. Insects associated with Nutgrass in Hawaii During this study in the summer of 1963, I came across the following insects associated with nutgrass. The aphid, Schizaphis cyperi, was collected on C. rotundus behind the laboratory in Hilo, Hawaii. Its occurrence seems to depend on the activity of ants which distribute the aphids from plant to plant. Most of the aphids were clustered on the basal stem portion of the plant. 3 Owing to a misidentification, Hawaiian records of S. mauritia prior to 1938 should apply to S. exempta (see Proceedings 1:75, 1938).

Vol. XVIII, No. 3, June, 1964 423 Rhopalosiphum rufiabdominalis (Sasaki)4 is a very common aphid found on roots of Cyperus esculentus in cane fields. Individuals of all sizes can be found on root clusters just beneath the tuber, and they are frequently attacked by coccinellid Tafvae. Their feeding often causes a general chlorotic condition of the plant. In one field, out of 5 plants of C. esculentus, 31 were infested with this aphid. Taeniothrips cyperi was common in the basal portion of the leaves in both C. rotundus and C. esculentus. When in large numbers, their feeding produces a scarring effect on the leaves which then show a purplish cast. Specimens of Dysmicoccus brevipes (Cockerell)5 were feeding on the tubers of both C. rotundus and C. esculentus. Saissetia nigra (Nietner)5 was found on leaves of many C. rotundus plants at Hilo during July. Several specimens of Chorizococcus (=Trionymus) rostellum (Hoke)5 were discovered on the roots of C. esculentus at Hilo, also during July. LITERATURE CITED Beardsley, J. W. 1963. Billbugs on Nutgrass. Coop. Econ. Insect Rept. USDA. 13(1) :186. Bianchi, F. A. 1945. Notes on Hawaiian Thysanoptera with Description of a New Species. Proc. Hawaiian Ent. Soc. 12(2):279-286. Ghosh, C. C. 1921. Supplementary Observations on Borers in Sugarcane, Rice, etc. Rept. Proc. Fourth Entomological Meeting at Pusa, 15-136. Marshall, Guy A. K. 1928. New Injurious Curculionidae. Bull. Ent, Research 18(3): 257-266, 5 figs. Meyrick, E. 199. Descriptions of Indian Micro-Lepidoptera. X Eucosmidae. Jour. Bombay Nat. Hist. Soc. XIX, 3:582-67. Pemberton, C. E. 1948. Nutgrass Insects. Hawaiian Planters' Rec. 52(l):78-79- Swezey, O. H. 1926. Recent Introductions of Beneficial Insects in Hawaii. Jour. Econ. Ent. 19(5):7l4-72. Swezey, O. H. 1927-28. Entomology. Report Comm. Expt. Sta. Hawaiian Sugar Planters Association. 19 and 16-17. Swezey, O. H. 1929. Nutgrass Borer {Bactratruculenta). Report Comm. Expt. Sta. Hawaiian Sugar Planters' Association. 18-19. Swezey, O. H. 1931. Nutgrass Borer {Bactratruculenta) Proc. of the Fiftieth Annual Meeting of the Hawaiian Sugar Planters' Assn. p. 427. Swezey, O. H. 1931a. Proc. of the Fiftieth Annual Meeting of the Hawaiian Sugar Planters' Association, p. 125. Swezey, O. H. 1935. Note. Proc. Hawaiian Ent. Soc. 9(1):32. Swezey, O. H. 1946. Note. Ibid. 12(3):475. Swezey, O. H. and Pemberton, C. E. 1931. Nutgrass Insects. Report Comm. Expt. Sta., Hawaiian Sugar Planters' Assn. p. 23. Williams, Francis X. 1931. Handbook of the Insects and other Invertebrates of Hawaiian Sugar Cane Fields, p. 4. Hawaiian Sugar Planters' Assoc, Honolulu. Zimmerman, E. C. 1936. Note. Proc. Hawaiian Ent. Soc. 9(2): 116. Zimmerman, E. C. 1948. Insects of Hawaii 2:414, 445. Univ. of Hawaii Press, Honolulu. Zimmerman, E. C. 1948a. Insects of Hawaii 5:1. Univ. of Hawaii Press, Honolulu. Zimmerman, E. C. 1958. Insects of Hawaii 7:35, 336. Univ. of Hawaii Press, Honolulu. 4 Determined by L. M. Russell, Entomology Research Division, Beltsville, Maryland. 5 Determined by J. W. Beardsley, University of Hawaii, Honolulu.